End-Maastrichtian tetrapod tracks from Mexico

Theropod, avian, pterosaur, and arthropod tracks from the uppermost

Cretaceous Las Encinas Formation, Coahuila, northeastern Mexico,
and their significance for the end-Cretaceous mass extinction

Wolfgang Stinnesbeck1,†, Eberhard Frey2, Belinda Espinoza-Chávez3, Patrick Zell4, José Flores-Ventura5,
Héctor E. Rivera-Sylva5, Arturo H. González-González6, José M. Padilla Gutierrez7, and Francisco J. Vega7
1
Institut für Geowissenschaften, Universität Heidelberg, Im Neuenheimer Feld 234, 69120 Heidelberg, Germany
2
State Museum of Natural History Karlsruhe, Erbprinzenstraße 14, 76133 Karlsruhe, Germany
3
Benemérita Escuela Normal de Coahuila, Calzada de los Maestros s/n, Saltillo, C.P. 25000, Mexico
4
Hessisches Landesmuseum Darmstadt, Friedensplatz 1, 64283 Darmstadt, Germany
5
Santa Engracia 257, Fracc. Santa Elena, Saltillo, Coahuila C.P. 25015, Mexico
6
Museo del Desierto, Carlos Abendrop Dávila No. 3745, Parque Las Maravillas, Saltillo, Coahuila C.P. 25015, Mexico
7
Instituto de Geología, Universidad Nacional Autónoma de México, Ciudad Universitaria, Mexico D.F. 04510, Mexico

ABSTRACT
Two unique localities that combine an un-
usual diversity of avian, pterosaurian, and
dinosaur tracks as well as trails of arthropods
were recently discovered by us in upper­
most Maastrichtian siliciclastic sediments of
the Las Encinas Formation in the Mexican
state of Coahuila, ~40 km north of Saltillo.
The trackway assemblages at Amargos and
­Rancho San Francisco were produced by at
least six different types of birds, while track-
ways of azhdarchoid pterosaurs are rare.
Only a single footprint was produced by a
nonavian theropod. A diverse ichnofauna of
arthropod traces is also present in a differ-
ent facies. The tetrapod trackway assemblage
was deposited during the very latest Maas-
trichtian, as indicated by an up to 2.5-m-
thick unit with abundant smectite spherules
attributed to the Chicxulub impact less than
8.5 m stratigraphically up section at Amar-
gos. Sphenodiscus pleurisepta is the last am-
monite at Amargos and may have crossed the
Cretaceous-Paleogene boundary.
INTRODUCTION
There has long been a fervent debate whether
the end-Cretaceous mass extinction was cata-
strophic, caused by the impact of a large bolide
(comet or asteroid) at Chicxulub into the Yuca-
tán Peninsula, Mexico, or gradual, caused by
climatic and sea-level changes triggered by the
Deccan volcanism (e.g., Schulte et al., 2009;
†
wolfgang​.stinnesbeck@​geow​.uni​-heidelberg​.de
Archi­bald, 2014; Renne et al., 2015; Keller et al.,
2016). Brusatte et al. (2015) suggested that the
abruptness of dinosaur extinction is the key to
advocate for the bolide impact, but the coarse-
ness of the fossil record makes testing the effects
of this hypothesis difficult (Archibald, 2014). To
date, few localities anywhere in the world host
nonavian dinosaurs in uppermost Maastrichtian
strata (e.g., France—Galbrun, 1997; Spain—
López-Martínez et al., 2001; North America—
Campione and Evans, 2011; Archibald, 2014). It
is therefore not unequivocally known whether or
not there was a global long-term decline of the
nonavian dinosaur diversity prior to their final
extinction at the end of the Cretaceous. Recently,
Sakamoto et al. (2016) argued for a long-term
decline in the diversity of dinosaurs, possibly
caused by global geotectonic phenomena (e.g.,
the breakup of Gondwana and Laurasia, intense
prolonged volcanism) and resulting fluctuations
in climate and sea level.
In contrast to nonavian dinosaurs, avian dino­
saurs and most other tetrapods such as squa-
mates, turtles, and crocodilians, as well as most
invertebrate groups, survived the Cretaceous-
Paleogene boundary (e.g., MacLeod et al., 1997;
Novacek, 1999; Gelfo and Pascual, 2001; Lutz,
2005; Evans and Klembara, 2005; Goin et al.,
2006; Longrich et al., 2011). This selectiveness
of extinction is a strong argument against a sud-
den impact-caused extinction. Brusatte et al.
(2015) suggested that the decline of large-bodied
herbivores in latest Cretaceous dinosaur assem-
blages of North America might have been caused
by a breakdown of the nutrient chain, which
would have made megavertebrate communities
more susceptible to cascading extinctions. Part
of this restructuring refers to the rise and diver-
sification of birds since the Early Cretaceous, al-
though the physical evidence for the abundance
and diversity of this group during the latest Cre-
taceous is presently rare (Longrich et al., 2011;
Jetz et al., 2012; Feduccia, 2014).
The search for answers regarding the timing,
causes, and course of the Cretaceous-Paleogene
boundary extinction has increasingly focused on
investigations concerning the late Maastrichtian
climatic changes and the dramatic oscillations
of sea level resulting in high-stress biotic envi-
ronments (e.g., Keller et al., 2016).
A complete, expanded, and fossil-rich sedi-
ment sequence covering the latest Maastrich-
tian to Cretaceous-Paleogene boundary into the
earliest Paleogene was recently discovered by
us in two localities in the eastern Parras Basin
(Difunta Group) of Coahuila, northeastern Mex-
ico. The investigated successions at San Juan
Amargos (Amargos in the following text) near
Paredón, and at a rancho named San Francisco
18 km to the west (Fig. 1) were deposited in a
marginally marine deltaic environment (Gold-
hammer and Johnson, 2001; Soegaard et al.,
2003) during about the last 100–200 k.y. of the
Maastrichtian (Vega et al., 2013). They present
ideal conditions in which to investigate pre–
Cretaceous-Paleogene changes in the vertebrate
and invertebrate communities in the region, in-
cluding the extinction of ammonites, pterosaurs,
and dinosaurs, and the diversification of birds at
that time.
We here document the tetrapod and inver-
tebrate trackways and faunal assemblages of
the new Amargos and San Francisco sites and
also discuss the paleoecological and geological

GSA Bulletin; Month/Month 2016; v. 128; no. X/X; p. 1–18; doi: 10.1130/B31554.1; 14 figures.; published online XX Month 2016.

For permission to copy, contact editing@geosociety.org

Geological Society of America Bulletin, v. 1XX, no. XX/XX 1
© 2016 Geological Society of America
 Stinnesbeck et al.

Coahuila
101°15′ 100°45′ 100°15′
A
57
Nuevo León Guadalupe 53

Anhelo
N San Juan Amargos

26°00′
W E San Francisco
S Paredón
114
MONTERREY
40 57
Hipólito
LA site

25°30′
General Cepeda SALTILLO

ila
hu
oa
eón

C
Nuevo L
30 km 54

Shallow, nearshore medium- Moderately deep marine Nonmarine to marginal-

Land p
to fine-grained clastics calcareous shale, siltstone marine coarse clastics
LLANO G rou
Outer-ramp lime Deep-marine, off-ramp UPLIFT rro
Study sites; San Juan Amargos: N25°56.502′/W101°01.392′
mudstone and shale lime mudstone and shale a va
San Francisco: N25°48.255′/W101°09.213′
N

B
e
al
Sh
ro
v ar
MAASTRICHTIAN Laredo Na
FORELAND
LA POPA
BASIN
Difunta
Difu Group
Gro nta
up Monterrey
ale

PARR
ez Sh

BASINAS Saltillo
Mend

MAASTRICHTIAN
UPLIFT
MAASTRICHTIAN
SHORELINE

200 km

Figure 1. Geographical and geological setting. (A) Geographic overview, where the sampling sites are marked with stars (map modified
after INEGI, 2015). LA site—Las Águilas. (B) Late Cretaceous (Campanian–Maastrichtian) paleogeography, following Goldhammer
and Johnson (2001). The sampling sites are marked with stars. Sediment transport was via rivers from the northwest, feeding an exten-
sive deltaic system in Coahuila and northwestern Nuevo León, known as the Difunta Group. Note that coastlines of the Difunta deltaic
complex prograded from west to east. To the east, in Nuevo León and Tamaulipas States, the Mendez Formation is coeval to the Difunta
Group and was deposited under open-marine conditions. North of the La Popa Basin, extensive paralic coal swamp systems developed
in the region of northern Coahuila. A NW-SE−extending fold-and-thrust belt is indicated by a dashed line with black triangles. Inset:
Paleogeographic map of North America during the Late Cretaceous (Campanian–Maastrichtian; from Blakey, 2015).

2 Geological Society of America Bulletin, v. 1XX, no. XX/XX

 End-Maastrichtian tetrapod tracks from Mexico

circumstances that caused the concentration of
footprints in the area. Based on the paleodiver-
sity discovered by us in the 70-m-thick (Amar­
gos) and 15-m-thick (San Francisco) sections
of latest Maastrichtian age and reaching up to
and across the Cretaceous-Paleogene bound-
ary, as well as a wide set of sedimentological
characteristics, we reconstruct a hitherto un-
known subtropical latest Cretaceous ecosystem
at the southernmost margin of the North Ameri-
can continent, only a few tens to hundreds of
thousands of years (k.y.) prior to the Chicxu-
lub impact.
GEOLOGICAL BACKGROUND
During the Late Cretaceous and Paleogene, a
large foreland basin developed north of the ris-
ing Sierra Madre Oriental in an extended area of
the Mexican states of Coahuila and Nuevo León
(Lawton et al., 2009). The basin was caused by
the subduction of the Farallon plate at the west-
1975; Sohl et al., 1991). The cyclical depo­si­tion
of alternating prodelta, delta front, delta plain,
and fluvial sediments was first subdivided into
lithostratigraphic units, or formations, by Mc-
Bride et al. (1974). To date, the Difunta Group
is known to extend stratigraphically from the
Upper Campanian to Eocene (e.g., Vega et al.,
1989, 2007; Soegaard et al., 2003; Ifrim et al.,
2010, 2015; Fig. 2).
Assignation of formations (e.g., Las Encinas
and Rancho Nuevo) to the Paleogene (Mid-
way stage) is based on the occurrences of the
nautiloids Cimomia haltomi (Aldrich, 1931)
and Hercoglossa sp., but also on the presence
of venericardiid bivalves. These ages were
first documented in Coahuila by Murray et al.
(1959) and subsequently confirmed by Hassel-
tine (1968), Wolleben (1977), and Vega et al.
(2007), among other authors. Ostreids were
Eustatic sea level curves,
3rd order cycles and magneto-
Age
also reported to be biostratigraphically signifi-
cant (e.g., McBride et al., 1974; Perrilliat and
Vega, 1993, 2003; Vega et al., 1999, 2007). The
Cretaceous-Paleogene boundary is placed in the
Las Encinas Formation, based on the first ap-
pearance of the nautilids mentioned above but
also on the presence, in the area north of Mon-
terrey (La Popa), of reworked Chicxulub impact
ejecta (Lawton et al., 2005; Schulte et al., 2012).
Tetrapod Remains from the Difunta Group
The entire area, extending from eastern Chi-
huahua to northern Nuevo León, is extremely
rich in well-preserved and diverse tetrapod re-
mains. The assemblages are mostly late Campa-
nian in age and include turtles and crocodilians,
among them eusuchians including the massive
alligatoroid Deinosuchus (Rivera-Silva et al.,
Correlation of formations
SC

ern margin of the North American plate, as well stratigraphy (Haq et al., 1987) of the Difunta Group
as by rapid subsidence related to the Late Cre- 250 200 150 100 50 0m
taceous Laramide orogeny (Weidie and Murray, Pal 1.2 DS

C30C29 C29
6 5
1967; McBride et al., 1974, 1975; Soegaard
Las Encinas Potrerillos

SC 3
et al., 2003; Lawton et al., 2009). North of the
late Maastrichtian

Sierra Madre Oriental mountain range, between 1.1 Cerro Grande 4

the cities of Saltillo and Monterrey, sediment
accumulated in this foreland from Campanian 3
to Paleogene times. This sediment sequence is Imagenes
known as the Difunta Group (Figs. 1A and 1B;

SC 2
McBride et al., 1974, 1975; Vega et al., 1989, 4.5 2
2007; Sohl et al., 1991; Soegaard et al., 2003; Cañon del Tule
C31

Stinnesbeck and Frey, 2014).

The Difunta Group represents shallow coastal
Muerto
and deltaic sediments and displays evidence of 1
coastlines of more than 500 km length (Figs.
early Maastrichtian

1A and 1B). The sediment sequence reaches Cerro Huerta

6000 m thickness or more proximal to the ­Sierra Cerro del Pueblo

SC 1
C32

Madre Oriental, but thicknesses gradually de-

crease to the north. In the Rio Grande area, 4.4
some 400–500 km north of Saltillo, their total
thickness is only 100–200 m (Cooper, 1970, Parras Shale
1971; Goldhammer et al., 1991). The Difunta
Group and coeval units in eastern Chihuahua
C33

Predominant lithologies:
and northern Coahuila States were deposited
Camp.

under deltaic conditions, representing marsh,
lagoonal, and eulittoral to shallow-marine envi-
ronments (Hopkins, 1965; Weidie et al., 1972;
Lehman, 1982).
The Difunta Group delta was fed by a river
system located in Chihuahua and western Coa-
huila and opening into the ancient Gulf of Mex-
ico. Progradation of the delta front gradually
moved the coastlines toward the east. During
the Campanian, coastlines reached the area of
Saltillo, and, during the Maastrichtian, the coast-
lines prograded further toward Monterrey (Figs.
1A and 1B; Weidie et al., 1972; McBride et al.,
sandstone siltstone
mudstone
4.3
Figure 2. Tentative correlation of the Difunta Group using bio-, magneto-, and chrono­
stratigraphic data, redrawn from Ifrim et al. (2010). C—magnetic chron. This correla-
tion is based on five stratigraphic marker horizons indicated by asterisks: 1—C32-C31
boundary, considered as base of the Maastrichtian (Eberth et al., 2004); 2—Lower
Cretaceous planktic foraminiferal zone (CF) CF 5 at La Parra; 3—age of 66.2 Ma for
the Imagenes Formation (Lawton et al., 2009); 4—sea-level lowstand at the base of
CF 3; 5—Cretaceous-Paleogene boundary within the Delgado Sandstone Member (DS)
in the La Popa Basin (Lawton et al., 2001), also present at San Juan Amargos; 6—Cre-
taceous-Paleogene boundary in the upper Las Encinas Formation (Vega et al., 2007).
SC—sedimentary cycles from Soegaard et al. (2003); Pal—Paleogene.

Geological Society of America Bulletin, v. 1XX, no. XX/XX 3


2011a), mosasaurs (Buchy et al., 2005), and
abundant and diverse dinosaur remains (Kirk-
land et al., 2006; Rivera-Sylva and Espinosa-
Chávez, 2006; Rivera-Sylva et al., 2006, 2009,
2011b, 2014, 2016; Gates et al., 2007; Ramírez-
Velasco et al., 2014; Vogt et al., 2016).
Dinosaur-bearing localities are known from
the Parras Basin at Las Águilas (LA site in
Fig. 1), near the hamlet of Porvenir de Jalpa,
and Rincón Colorado near the town of General
Cepeda. The deposits assigned to the Cerro del
Pueblo Formation are late Campanian in age
and display a rhythmical sequence of sandstones
and siltstones deposited under littoral conditions
of changing salinity, as is indicated by oyster
banks, and abundant non-ostrean bivalves and
gastropods, but also sharks, mosasauroids, and
dyrosaurid crocodilians and the shallow-water
ammonite Sphenodiscus. Freshwater to brack-
ish conditions are indicated by gastropods,
characean oogonids, vascular plants, shell frag-
ments of clemmydid and trionychid turtles,
small eusuchian crocodilians and dinosaurs,
among them abundant hadrosaurids and, more
rarely, lambeosaurids, ceratopsians, and non­
avian theropods (e.g., Eberth et al., 2004; Rivera-
Sylva et al., 2006, 2011b, 2014; Rodríguez-de la
Rosa, 2007; Meyer et al., 2008; Perrilliat et al.,
2008; Vogt et al., 2016). Vertebrate remains in-
cluding pterosaur trackways have been described
from the El Pelillal area and were also dated to
the late Campanian (Rodríguez-de la Rosa and
Cevallos-Ferriz, 1998; Rodríguez-de la Rosa,
2003). However, it was impossible to find this
locality based on the data annotated in the pub-
lications (Stinnesbeck and Frey, personal obs.,
2015). Up section, Maastrichtian units of the Di-
funta Group are much less well studied in terms
of vertebrate assemblages, and their dinosaurs,
or other vertebrate remains, are largely unknown
(Rivera-Sylva et al., 2014).
Little is known about tetrapods from the Up-
per Maastrichtian units of the Difunta Group
(e.g., the Las Encinas Formation), because the
fossil record is sparse and cannot be properly
dated. Recently, Schulte at al. (2012) reported
on an isolated dinosaur bone in a sandstone
unit containing Chicxulub impact spherules in
the La Popa Basin near Monterrey, but the only
material illustrated was attributed by the authors
to a mosasaur (Schulte et al., 2012, their figs.
9A and 9B). It is therefore impossible to verify
whether or not the bone referred to is really
dino­saurian. In addition, the bone was described
as “abraded” and may therefore have been re-
worked from an older stratigraphic unit.
Even though the fossil record of late Maas-
trichtian dinosaurs in Mexico is presently incon-
clusive, the Difunta Group delta must have been
rich in life and certainly attracted pterosaurs and
4 Geological Society of America Bulletin, v. 1XX, no. XX/XX
Stinnesbeck et al.
birds in their search for food and freshwater.
The rivers, their banks, and the expanded coastal
sand and mud plains provided all they needed
for survival: Small epibenthic and endobenthic
animals and fishes must have been abundant.
Carcasses of tetrapods were likely washed up
in small bays and attracted scavengers of all
kinds, including aerial ones. The hollow, brittle
bones of pterosaurs and birds are rapidly eroded
by floods and sediment flow, and therefore only
small, mostly undiagnostic fragments of ptero-
saur long bones have been found in the Parras
Basin. Some of them were tentatively referred
to Ornithocheiroidea indet., a diverse group
of pterosaurs that was widespread throughout
the Late Cretaceous (Barrett et al., 2008). The
identi­fication is based on the oval cross section
of a fragmentary possible distal wing finger
phalanx (Rodríguez-de la Rosa, 1996). How-
ever, our reexamination of the material housed
in the Museo del Desierto, Saltillo, could not
positively confirm this interpretation. Birds are
yet undocumented from the Difunta Group.
MATERIAL AND METHODS
Localities
In 2013, two of us (Espinoza-Chávez, Flores-
Ventura) discovered the two sites documented
here (Amargos, San Francisco). San Juan Amar­
gos, at ~10.7 km SW of Paredón (25°56.502′N,
101°01.392′W), is well exposed, and it con-
tains a Chicxulub impact spherule unit that un-
equivocally assigns the top of this section to the
Cretaceous-Paleogene boundary. The Amargos
section is here used as a reference to correlate
this sediment sequence and its near–Cretaceous-
Paleo­gene boundary spherule unit to a section lo-
cated ~18 km west of Amargos, at San Francisco
(25°48.255′N, 101°09.213′W). In this latter lo-
cality, the Chicxulub spherule unit is absent, but
the presence of invertebrate fossils, among them
the Paleocene turkostreid Gorizdrella gorizdroae
Vyalov, 1937, in strata overlying Maastrichtian
faunal assemblages, including the tetrapod track-
way unit, argue in favor of an expanded Creta-
ceous-Paleogene boundary transition.
Field Work and Sections
Field work for the present study was carried
out at Amargos in April 2014, September 2015,
and March 2016. The San Francisco site was
investigated in April 2014 and March 2016. An
~78-m-thick section of the uppermost Maas-
trichtian–Lower Paleogene Las Encinas Forma-
tion (Fig. 3) was measured and sampled to strati-
graphically assign the vertebrate track-bearing
layers and invertebrate remains and to locate the
Cretaceous-Paleogene boundary in the Amar­
gos section. The layer-by-layer investigation
included details on vertebrate track occurrence,
invertebrate and plant fossils, and sediment struc-
tures, as well as records of facies and taphon­
omy (Fig. 4). Significant fossils were collected.
Detailed data on the stratigraphic position, size
class, number of individuals, preservation state,
etc., were also collected on fragmentary or al-
ready registered specimens, but the majority of
these specimens remained in the field.
Specimens
All collected fossils are housed in the Colec-
ción de la Escuela Normal Superior de Coahuila
and were also given numbers in the Colección
de Paleontología de Coahuila (CPC), at the
Museo del Desierto, Saltillo, Coahuila, Mexico,
including the fossils illustrated here.
Photography
Photographs were made with an Olympus E
620 SLR digital camera with a Zuiko Digital
14–42 mm, 1:3.5–5.6 lens and a Zuiko Digi-
tal Macro 35 mm, 1:3.5 lens. Low-angle light
photography was carried out with an infrared
controlled Olympus Electronic Flash FL 36R.
The photographs were traced with CorelDraw
X6. The drawings were designed with the
same program.
RESULTS
San Juan Amargos (Amargos)
At Amargos, uppermost Maastrichtian gray
mudstones of the Las Encinas Formation include
crustaceans, and abundant mollusk (e.g., the
ammonite Sphenodiscus pleurisepta) and turtle
remains, but also abundant invertebrate tracks
and burrows, plant debris containing wood of
palms and conifers, diverse and abundant avian
tracks, and rare monotypic pterosaurian tracks.
Except for the crustaceans documented by Vega
et al. (2013; e.g., Ophthalmoplax brasiliana,
Costacopluma grayi, Linuparus sp., callianas-
soid remains), the San Juan Amargos fossil as-
semblage is undocumented to date.
Vega et al. (2013) assigned the Amargos sec-
tion to the Las Encinas Formation, which in the
Parras Basin crosses the Cretaceous-Paleogene
boundary (e.g., Murray et al., 1960; Weidie
and Murray, 1967; Hasseltine, 1968). The au-
thors suggested that the section contained the
last 500 k.y. of the Cretaceous and the Creta-
ceous-Paleogene boundary. This interpretation
is supported by detrital zircon ages of 66.2 Ma
(minimum age) for the Las Imágenes Formation
 End-Maastrichtian tetrapod tracks from Mexico

A W E

2m

B C

0.5 m 0.5 m

D E F
2
1

0.1 m 2m 0.4 m

Figure 3. Outcrop photographs of the uppermost Maastrichtian–Lower Paleogene Las Encinas Formation at San Juan Amargos (A–D) and
San Francisco (E–F). (A) Basal-most siltstone (meters 2–11) rich in marine fossils at Amargos. (B) Channelized sandstone rich in fossil wood
(meters 29–31 of the Amargos section). (C) Near−Cretaceous-Paleogene boundary channelized microconglomerate and sandstone unit con-
taining Chicxulub-impact spherules (meters 70–72 at Amargos). The spherule-rich layer is at the base of the siliciclastic unit and is marked
with an arrow. (D) Detail of C: Microconglomerate and sandstone. (E) Upper part of section at Rancho San Francisco: 1—sandstone at
top of section, 2—claystone unit with Gorizdrella gorizdroae Vyalov, 1937. (F) Unit with rippled platy sandstone with abundant tetrapod
trackways at Rancho San Francisco. Meters refer to the lithostratigraphic sections illustrated in Figures 4 and 8.

underlying the Las Encinas Formation (Lawton
et al., 2009) and by the presence of Paleocene
ostreid banks from the upper part of the Las
­Encinas Formation (Vega et al., 1999).
Stratigraphic Sequence (Fig. 4)
The base of the section, assigned to the basal
Las Encinas Formation by Vega et al. (2013),
consists of a 6-m-thick bioturbated (e.g., Chon-
drites) gray siltstone with crustaceans (e.g.,
Costacopluma grayi), abundant bivalves (e.g.,
cardiids, oysters), gastropods (e.g., cerathiids),
the ammonite Sp. pleurisepta, and turtle, chon-
drichthyan, and actinopterygian remains (e.g.,
teeth of lamnoid sharks, toothplates of the saw
fish Schizorhiza). Up section, an interval of
~4 m of gray siltstone and claystone contains
an increasing number of yellow-buff–colored
cross-bedded and rippled sandstone units from
0.2 m to 1 m thick, which are channelized and
present undulated basal contacts. The top of
the sandstone units is bioturbated by Ophio-
morpha. The interval between 10 and 45 m
from the base of the section is increasingly rich
in sandstone, while intercalated gray siltstone
layers are reduced to a few tens of millimeters
to a maximum of 0.2 m thickness. Sandstones

Geological Society of America Bulletin, v. 1XX, no. XX/XX 5

 Stinnesbeck et al.

40 80
[m] change in the
oyster assemblage

Paleocene

Gorizdrella gorizdroae
K/Pg-channel
with spherules

Difunta Group – Las Encinas Formation

30 70

uppermost Maastrichtian

Flemingostrea
Difunta Group – Las Encinas Formation
uppermost Maastrichtian

20 60

microconglomerate sandstone

siltstone marl clay

cross wavy nodular
bedding ripples bedding
erosive mud
contact concretions clasts

f/m/c - fine/medium/coarse sst.

unexposed section fish
10 50
other marine
vertebrates crustaceans

ammonites oysters
other gastropods
bivalves
sponges vertebrate
tracks
other bivalve
tracks shell beds
bioturbation fragmented
wood fern

0 40
fm f mc

Figure 4. Uppermost Maastrichtian sediments and the Cretaceous-Paleogene (K/Pg) boundary transition at Amargos, Coahuila, showing
the lithostratigraphic distribution of important fossils and the stratigraphic position of vertebrate tracks. Tracks of insects and other arthro­
pods (e.g., sediment-feeding crustaceans) were detected on sandstone (sst) surfaces at meters 32 and 38 of the section. Vertebrate tracks were
discovered in a 50–100 mm thin layer at 62 m of the section, and thus only 8.5 m below the base of a channelized siliciclastic unit containing
abundant Chicxulub impact spherules. Note that the lower, fossil-rich portion of the sediment sequence and the Cretaceous-Paleogene-
boundary transition at San Juan Amargos are predominantly marine. The middle portion of the succession, between meters 18 and 45 of the
section, represents deltaic depositional environments, as indicated by the absence of marine-indicative faunal elements, the low fossil content,
and the presence of characteristic sedimentary structures (e.g., cross-bedding, ripple-bedding, tidal channels, desiccation cracks).

6 Geological Society of America Bulletin, v. 1XX, no. XX/XX


are cross-bedded and present rippled surfaces,
with mud cracks, algal mats, rare oysters,
abundant wood and twigs, and occasional bio-
turbation by Ophiomorpha. Some sandstone
units appear to be channelized or incised into
underlying silt. These structures likely repre-
_ A
__
_ F
_ _
E San Francisco section, Coahuila,
I (C) Left valve, specimen CPC-
_
M N
_ _
_
__
_ __
Geological Society of America Bulletin, v. 1XX, no. XX/XX 7
End-Maastrichtian tetrapod tracks from Mexico
sent ancient channel tideways in a delta surface
environment.
A unit with abundant tetrapod trackways
(avians, azhdarchoid pterosaurs) and oysters
(Fig. 5) was discovered at meter 61 in the sec-
tion. It is only ~0.2–0.3 m thick and is com-
B _ C D
_G
_
J
__
K L
_ _
posed of y­ ellow- to buff-colored platy sandstone
layers of a few tens of millimeters in thickness.
Locally, the sandstone preserves ripple marks.
Approximately 8.5 m of unfaulted section
separate the sandstone with tetrapod trackways
from a channelized unit of microconglomer-
H
Figure 5. (A–P) Gorizdrella goriz­
droae Vyalov, 1937, early Paleo-
cene, Las Encinas Formation.
(A–I) Specimens from Rancho
at 11–12 m of the section depicted
in Figure 8. (A) Left valve, speci-
men CPC-1879. (B) Left valve, de-
tail of hinge line, same specimen.
1880. (D) Left valve, inner view,
same specimen. (E, F) Left and
right valve views, specimen CPC-
1881. (G–I) Left, right, and lat-
eral views, specimen CPC-1882.
(J–P) Specimens from San Juan
Amargos section, Coahuila (for
stratigraphic position see Fig. 8).
(J) Left valve, specimen CPC-
O 1883. (K, L) Left valve and lat-
eral views, specimen CPC-1884.
(M) Agglomerate of left valves,
specimen CPC-1885. (N) Left
valve , s pe c ime n CPC- 1886.
(O) Left valve next to Chicxulub-
impact spherules (blue lines),
specimen CPC-1887. (P) Left
valve next to Chicxulub-impact
spherules (blue lines), specimen
CPC-1888. Scale bars = 10 mm.
P

ate and sandstone containing Chicxulub impact
spherules (Fig. 6). This interval is intermittently
exposed and consists of gray siltstone and ­yellow
sandstone. Oysters are abundant (Fig. 5), but
Sp. pleurisepta (Figs. 6 and 7), bryozoans, and
serpulids also occur. Oysters in this interval are
tentatively assigned to Flemingostrea, which is
a widely distributed genus in the Upper Maas-
trichtian sediments of the area (Wolleben, 1977;
Vega and Perrilliat, 1989; Perrilliat and Vega,
1993, 2003; Vega et al., 1999, 2007). However,
a significant change in the oyster assemblage is
observed ~0.2–0.5 m below the erosive base of
the Chicxulub spherule–bearing unit (Fig. 4).
In these sediments, another oyster taxon occurs
that is different from any Maastrichtian or Paleo-
gene species previously reported for the Difunta
Group (Fig. 5). The taxon is morphologically
identical to Gorizdrella gorizdroae Vyalov, 1937
from the early Paleogene of Central Asia (e.g.,
Afghanistan; Berizzi Quarto di Palo, 1970).
The near–Cretaceous-Paleogene boundary
unit with spherules continuously crops out over
a distance of 80 m in a W-E direction and clearly
represents an incised channelized deposit, with
a thickness of ~1.6 m at the easternmost and
1.8 m at the westernmost exposed margin, but
~2.3 m in the outcrop center (Fig. 6). The base
of the unit is erosive and consists of up to 11
distinct layers of amalgamated oligomict micro­
conglomerate with angular to subrounded
clasts, sandstone, and interlayered siltstone and
shale. These layers mostly lack internal struc-
tures, but they are locally cross-bedded, while
other beds are slightly graded. Oysters are
abundant in the basal unit and include G. goriz-
droae. They locally form shell hash of highly
fragmented small individuals, while other lay-
ers contain specimens larger than 60 mm that
are mostly unfragmented. Shark teeth, bone
fragments, well-preserved Sp. pleurisepta
specimens (Conrad, 1857), turritellid gastro-
pods, bivalves (e.g., cardiids), and wood frag-
ments are also abundant. Spherules are visible
by eye or hand lens in the basal layer of the
siliciclastic unit, and abundant (Fig. 6A; 20%),
but isolated rare spherule specimens were also
detected during thin section analysis in overly-
ing layers (Fig. 6D). The spherules are ~1 mm
in diameter and mostly rounded or elongated,
although crushed specimens also occur. They
are completely devitrified to smectite and mixed
into siliciclastic debris, but they mostly preserve
their original vesicular texture. Voids are filled
with blocky calcite, although broken voids are
commonly filled with matrix material. These
vesicular devitrified spherules provide a critical
link to the end-Cretaceous Chicxulub impact in
southern Mexico, as evidenced in deep-marine
Cretaceous-Paleogene boundary sites in the
8 Geological Society of America Bulletin, v. 1XX, no. XX/XX
Stinnesbeck et al.
Burgos Basin east and south of Monterrey and
elsewhere in the region (Keller et al., 2003a,b;
Schulte et al., 2010), but also from a similar
shallow channelized deposit with Chicxulub
spherules in the La Popa region north of Mon-
terrey (Schulte et al., 2012).
Sp. pleurisepta (Conrad, 1857) is the latest
ammonite appearance in northeastern Mex-
ico (Ifrim et al., 2005). At La Popa, it occurs
throughout the Maastrichtian up to 3 m below
the Cretaceous-Paleogene boundary (Lawton
et al., 2001). At Amargos, we collected several
specimens from the unit with reworked spher-
ules, up to ~0.6 m above the base of this deposit.
Our collection includes well-preserved com-
plete specimens of Sp. pleurisepta, even with
spherules in its body chamber (Figs. 6 and 7).
Rancho San Francisco
About 15 m of claystone, siltstone, and sand-
stone of the Las Encinas Formation were sam-
pled at Rancho San Francisco. Results of our
lithostratigraphic survey are illustrated in Fig-
ure 8. The base of the section, which is intermit-
tently exposed over a distance of at least 200 m
along the hillside, includes an ~0.5-m-thick,
yellow- to buff-colored sandstone unit with
common trackways of birds, abundant azhdar-
choid pterosaur manus and pes imprints, and a
single nonavian theropod footprint. Casts, likely
produced by burrowing annelids, are also abun-
dant on the upper surfaces of these sandstones.
The sandstone is mostly platy, but layers with
well-preserved oscillation ripples and o­ thers
with desiccation cracks also occur. Oysters
(Flemingostrea) are the only body fossils identi-
fied in the sandstone and under- and overlying
siltstone.
Sediments overlying the sandstone with tetra-
pod trackways consist of gray brittle claystone
and siltstone forming units of up to 2 m, and
rare intercalated gray to yellow sandstone layers
a few tens of millimeters to a maximum of 0.2 m
in thickness. Only the lowermost ~13 m were
measured in detail (Fig. 8). The section above
is massively overgrown and thus difficult to as-
sess. While sandstones are devoid of body fos-
sils, the mudstone contains rare oysters (Flemin-
gostrea). A claystone unit with abundant cardiid
bivalves (e.g., Venericardia) and G. gorizdroae
was identified at ~8 m above the tetrapod track-
way–bearing sandstone and thus in a relative
stratigraphic position similar to that at Amargos.
Up section, ~20 m above the top of the San
Francisco section measured here, an intensively
red-colored sandstone ~1.5 m thick still forms
part of the Encinas Formation. Vega et al. (1999)
reported Paleocene oyster banks from the upper
Las Encinas Formation, in marine sandstones
from localities near the San Francisco site.
Correlation between Amargos
and San Francisco
The interpretation of Amargos as a Creta-
ceous-Paleogene boundary locality is now sup-
ported by the identification of Chicxulub impact
ejecta (smectite spherules) in the section ~60 m
above layers containing well-preserved crusta-
ceans (Vega et al., 2013), and ~8.5 m above a
fine-grained sandstone containing trackways of
birds and pterosaurs (Fig. 8). In addition, a layer
with abundant small-sized turkostreoids here
assigned to the Asian G. gorizdroae was dis-
covered ~0.3 m below the erosional contact that
forms the base of the Chicxulub ejecta-bearing
unit. This taxon is also present at San Francisco,
~8 m above the tetrapod trackway unit, which
there includes a single dinosaur footprint as
well as abundant and diverse avian footprints
and trackways and rare manus and pes imprints
of azhdarchoid pterosaurs (Fig. 8). Although no
Chicxulub ejecta appear to be present at San
Francisco, the uppermost Maastrichtian unit
with tetrapod trackways and overlying unit with
G. gorizdroae, of likely early Paleocene age,
confine the position of the Cretaceous-Paleo-
gene boundary to an interval <8 m in maximum
thickness (Fig. 8).
Vertebrate Ichnofossils
We identified uppermost Maastrichtian tetra­
pod ichnofossils in well-defined and correlat-
able stratigraphic units at both Amargos and
San Francisco. The majority of these ichno­
fossils were produced by a diverse avifauna
consisting of, to date, five or six distinguishable
kinds of tracks that have not yet been diagnos-
tically investigated in detail. Here, we present
a preliminary analysis and identify track types
(for position in the section, see Figs. 4 and 8).
Preliminarily, we stick to the description of the
morphotypes and present one typical example
of each. A comparative analysis of the track
types (e.g., stride length and width analyses of
the trackways) is beyond the scope of this paper
but is in preparation, based on new material.
Aves (Figs. 9–11)
Morphotype 1 (Fig. 9A). The standard length
for morphotype 1 is ~65 mm, as is the width,
with stout toes and webbing at the base of the
toes (semipalmate). The pes is tetradactyl, with
digit I up to 50% of the length of the longest
digit II, which also is the stoutest of the digits.
The medial orientation of digits II and IV sug-
gests highly mobile toes, similar to extant ibises,
the feet of which much resemble the morphol-
ogy of morphotype 1. Morphotype 1 is the most
abundant in the assemblage.
 End-Maastrichtian tetrapod tracks from Mexico

Figure 6. Detailed sedimentary columns of the near−Cretaceous-Paleogene-boundary siliciclastic unit at San Juan Amargos and repre-
sentative thin sections. The channelized deposit is highlighted in gray, and the stratigraphic positions of thin sections within the sediment
sequence are indicated by letters A to E. (A) Bioclastic oligomict microconglomerate with sand and micritic matrix and abundant oysters
and rare other bivalves. This horizon contains spherules, which are pointed out by arrows. Spherules are devitrified, weathered, and
encased by an orange colored mineral, but their outline and vesicular interior texture (e.g., A3 , A4 ) are preserved. (B) Sandstone (~40%
siliciclastic debris) with a micritic calcareous matrix and fragmented oysters. (C) Oligomict microconglomerate with sandy matrix and
rare oyster and gastropod bioclasts. (D) Oligomict microconglomerate with a sandy matrix. Oyster, gastropod, and echinoderm bioclasts
are rare, and altered spherules are abundant. (E) Oligomict microconglomerate with rounded to subrounded limestone, chert and calci-
sphere clasts, large oyster fragments, and a silty to sandy matrix. Abbreviations: f, m, c—fine, medium, coarse, respectively.

Geological Society of America Bulletin, v. 1XX, no. XX/XX 9


A1
10 mm
B photograph of a second speci-
Morpohotype 2 (Fig. 9B). The standard
length of the tetradactyl footprint is ~65 mm. As
in morphotype 1, the length of digit I is half the
length of digit II, with the latter being the ­longest
digit. The width of the footprint measures
~50 mm. In contrast to morphotype 1, digits II
and IV are almost straight or even curved cau-
dally. There is no webbing trace visible between
the toes (apalmate). It is still possible that mor-
photype 2 is conspecific with morphotype 1, and
the lack of a webbing imprint is biased by the
material properties of the sediment.
Morphotype 3 (Fig. 10A). The tiny, strongly
asymmetrical and apalmate footprint is 30 mm
long and 25 mm wide. A metatarsal pad is visi­
ble in some of the prints, but the birds must
have walked with an elevated metatarsus. Digit
I is half as thick as the other three digits and a
little less than half as long as the longest digit
(digit II). The toe prints are all straight, suggest-
ing a rigidity of the interphalangeal joints. The
10 Geological Society of America Bulletin, v. 1XX, no. XX/XX
Stinnesbeck et al.
A2
10 mm
angle between digits II and III is the largest of
all interdigital angles. The morphology of the
pes resembles the foot of small extant galli-
form birds.
Morphotype 4 (Fig. 10B). The tridactyl foot-
print is ~35 mm long with stout digits, of which
the second is the shortest. Digits II and IV face
laterally and medially, respectively, and span
~60 mm, which makes the footprint about twice
as wide as it is long. The stoutness of the digit
traces in relation to those depicted in Figure
10A is probably a result of preservation.
Morphotype 5 (Fig. 11A). This morphotype
is represented by a very faint tridactyl foot-
print. The interdigital angles are ~30°, yielding
a footprint width of only 16 mm. The webbing
traces reach the tips of the straight toes. There
are no traces of claws beyond the margin of the
webbing. Extant tridactyl seagulls possess simi-
lar foot morphologies, but they have slightly
curved, laterally convex digits II and IV.
Figure 7. Highest occurrence
of the ammonite Sphenodiscus
pleurisepta (Conrad, 1857) at
San Juan Amargos. (A) Speci-
men CPC-1889, A1, lateral
view, and A2, longitudinal sec-
tion with sketch (to the right)
to illustrate the distribution of
spherules from the Chicxulub
impact (dark dots inside the
body chamber). (B) Outcrop
men (CPC-1890). Specimens
come from the lowermost bed
of the Cretaceous-Paleogene-
channel, level of thin section A
in Figure 6.
Morphotype 6 (Fig. 11B). This tetradactyl
apalmate morphotype is the largest of the as-
semblage, with a length of more than 85 mm
and a width of ~70 mm. The digits are ex-
tremely asymmetrical, and the angle between
digits II and III is twice as wide as the angle be-
tween digits III and IV, similar to morphotype 2.
The imprint of digit I is reduced to a subcircular
to elongate oval pit. Digit I is about one quarter
of the length of digit II, which is the longest of
all pedal digits. The lack of a metatarsal ­cushion
and imprints of the basal phalanges suggest a
digitigrade locomotion habit with an elevated
metatarsus. The overall morphology of the foot
closely resembles that of extant gruiform birds.
Nonavian Theropods (Fig. 12)
A single tridactyl footprint of a right pes,
observed at Rancho San Francisco, was pro-
duced by a nonavian theropod dinosaur, prob-
ably a tyrannosaurid (Fig. 12). It has a length
 End-Maastrichtian tetrapod tracks from Mexico

WSW ENE
15 kilometers

San Francisco San Juan Amargos microconglomerate

20
[m] sandstone

siltstone

clay erosive Figure 8. Correlation between

contact
Paleocene

the Rancho San Francisco and

cross wavy San Juan Amargos Cretaceous-

Gorizdrella gorizdroae
bedding ripples Paleogene (K/Pg) boundary
f/m - fine/medium sst. sections. Note that a channel-
Las Encinas Formation

unexposed section ized Chicxulub impact ejecta−

bearing microconglomeratic
K/Pg-channel with spherules unit is present at Amargos but
absent at San Francisco. Cor-
unzoned

relation is achieved by the first

10 Gorizdrella appearance of the turkostreid
gorizdroae oyster Gorizdrella gorizdroae in
other marine the two sections, and the occur-
fish vertebrates rence of vertebrate trackways
(avians, pterosaurs, nonavian
uppermost Maastrichtian

Flemingostrea
ammonites oysters
therosaurs) at ~8 m below the
other first appearance datum (FAD)
gastropods
bivalves
of G. gorizdroae.
vertebrate bivalve
Flemingostrea shell beds
tracks
tet
rap
od fragmented bioturbation
ract
aves ks
pterosaurs
non-avian theropods
0
fm fm

of ~200 mm and is twice as long as wide, with
digit III being the longest and most massive
one. A large, rounded triangular metatarsal pad
forms a little less than one third of the entire
length of the preserved footprint. The toes are
almost parallel each other. The tips, including
the claw marks, of all toes are missing. Between
the toes and around the metatarsal pad, the sedi-
ment has been pushed up, suggesting the pres-
ence of a superficial layer of a muddy substrate
with a thickness of ~50 mm to a maximum of
100 mm. Below this layer, the substrate must
have been consolidated because the pes print
is shallow, and its penetration into the sediment
was stopped, even during the rolling phase. The
indistinct outline of the digital imprints without
any traces of digital pads suggests that this foot-
print represents an undertrack. Because of the
unclear preservation of the distal elements of the
digits, an identification beyond Tyranno­sauridae
appears problematic. Ornithomimosaurs show
diverging digits (Lockley et al., 2011), and
dromaeosaurs produce didactyl tracks (e.g.,
Lockley et al., 2016), and so these are also pre-
cluded as potential track makers.
Pterosauria (Fig. 13)
Manus (e.g., Fig. 13A) and pes prints (e.g.,
Fig. 13B) of pterosaurs are extremely rare in
these deposits and are mostly represented by
single imprints. While most of the manus prints
are undiagnostic (Fig. 13A), the pes prints show
a metatarsal area that has more than twice the
length of the webbed digital area (Fig. 13B).
Such a pes/metatarsus ratio is documented for
Brazilian azhdarchoid pterosaurs (Fig. 13C;
Frey et al., 2003). The longest pedal imprints
detected in the Mexican localities reach a length
of ~160 mm, which would suggest a wingspan
of ~4.5 m. One single slab exposes an isolated
130-mm-long tridactyl manus print of a likely
azhdarchoid pterosaur, which is inferred from
the short and stubby digits (Fig. 13A). The size
of the manus print suggests that the animal had a
wingspan exceeding 8 m. The producer may well
have been the size of a Quetzal­coatlus ­northropi,
which is known from the Upper Maastrichtian
Javelina Formation of Texas (Lawson, 1975).
The largest Cretaceous pterosaur pes tracks are
presently known from Korea and reach a length
of 340 mm (Hwang et al., 2002).
Invertebrate Tracks (Fig. 14)
Abundant invertebrate ichnofossils were
identified on the upper surfaces of yellow-buff
fine-grained sandstone in the interval between
meter 37 and 38 of the Amargos section. These
sandstone layers show massive burrows of
Ophiomorpha (Fig. 14E). On fine silty surfaces
of yellow tan sandstone, traces of epibenthic in-
vertebrates are abundant. A thorough investiga-
tion is pending, but the following morphotypes
are presently distinguished:
Morphotype 1 (Fig. 14A). This trackway
consists of a pair of scratch furrows that resem-
ble Cruziana D’Orbigny (1842); however, the
width of these tracks never exceeds 10 mm, and
the scratch marks of the legs are more oblique.
They were likely produced by small notostracan
crustaceans (e.g., Triops). However, the track
maker needs to be verified.
Morphotype 2 (Fig. 14B). This type consists
of a meshwork of furrows up to a diameter of
~8 mm or less without any further structure.
These traces might be referred to annelids, tiny
bivalves, or gastropods moving on the surface,
which, according to their abundance, may have

Geological Society of America Bulletin, v. 1XX, no. XX/XX 11

 Stinnesbeck et al.

III IV 1975; Soegaard et al., 2003; Vega et al., 2013).

webbing
ca. 70°
II
ca. 90°
A 30 mm
III
ca. 60°
ca. 75°
IV
B 30 mm
represented an exhaustive food source (e.g., for
birds). A closer investigation is pending.
Morphotype 3 (Fig. 14C). Transverse oval
opposing imprints aligned with a medial furrow
probably represent an enigmatic invertebrate
trace from the site. This track may have been
produced by a crustacean with ventrally fac-
ing uropods.
Morphotype 4 (Fig. 14D). The character-
istic features for this type of track are punc-
ture marks in the sediment. The width of the
trails is no more than 6 mm. The track maker
may have been an insect, a nonarachnid che-
licerate (e.g., a small scorpion), or an isopod
crustacean.
ca. 60°
Figure 9. Avian footprint types
from the Rancho San Fran-
cisco site: (A) morphotype 1,
I and (B) morphotype 2. The
interpretive line drawing is to
II
the right of the original photo-
graph for the selected footprint.
Roman numbers refer to digit
numbers of the pes.
ca. 130°
I
Morphotype 5 (Fig. 14E). This morphotype
has been identified as Ophiomorpha.
DISCUSSION
Paleoecological Setting
The uppermost Maastrichtian to lowermost
Paleogene Encinas Formation constituted a
system of slow-flowing shallow rivers and ox-
bows deposited in a coastal delta plain that was
dominated by small, low-sinuosity channels,
levees, and lagoons, with settings of intermit-
tent fresh and brackish water as well as shallow-
marine conditions (e.g., McBride et al., 1974,
Shallow-­marine environments are indicated by
the abundance of Sp. pleurisepta specimens in
repeated units at Amargos (Fig. 4). The taxon is a
common faunal element in the Difunta Group of
northern Mexico (Ifrim and Stinnesbeck, 2010)
and an excellent facies indicator for shallow-­
marine to proximal nearshore environments
(­Ifrim et al., 2005; Ifrim and Stinnesbeck, 2010).
On the other hand, the sandstone units of the Las
Encinas Formation containing arthropod and
tetrapod trackways are characterized by ripple
marks and patchy remnants of bacterial mats that
show desiccation cracks and flakes. Likely, this
sediment was accumulated in extremely shallow
waters including intermittent subaerial condi-
tions. These extended mud banks, full of micro-
bial life and bioturbation, provided the potential
for exquisite preservation of tracks of all kinds.
The abundance of life is indicated by the
wealth of invertebrate fossils (e.g., gastropods,
bivalves, crustaceans) and remains of fishes and
turtles in the basal siltstone unit of Amargos,
but also by the wealth of invertebrate trackways
attributed to small arthropods and annelids.
Therefore, the end-Maastrichtian water-rich
open plains in the area represented abundant
epibenthic and endobenthic food sources. This
is also suggested by the abundance of vertebrate
trackways, which in places even accumulate to
trampling horizons.
The diversity of the trace fossils evidently
depended on the presence of this mud-plain en-
vironment and its selective effects concerning
the fossil record of ichnotaxa, especially with
respect to aerial taxa.

II IV
A A ca. 30° webbing

ca. 85° III

III ca. 80°

I
ca. 30°
ca. 45°

II 10 mm
IV
10 mm
IV

B
III
B
II ca. 70°
ca. 60°
webbing
I
III
ca. 55°
ca. 120°
ca. 45°
IV

30 mm 30 mm
II

Figure 10. Avian footprint types from the Rancho San Francisco Figure 11. Avian footprint types from the San Francisco (A) and
site: (A) morphotype 3, and (B) morphotype 4. The interpretive line Amargos (B) sites: (A) morphotype 5, and (B) morphotype 6. The
drawing is to the right of the original photograph for the selected interpretive line drawing in A and B is to the right of the original
footprint. Roman numbers refer to digit numbers of the pes. photograph of the footprints.

12 Geological Society of America Bulletin, v. 1XX, no. XX/XX


A 2015). This pattern of erosion and reworking of
B the conglomerate forming the base of the Upper
III
damaged areas
pushed-up material
30 mm
Figure 12. Theropod footprint from Rancho San Francisco:
(A) origi­nal photograph of the footprint, and (B) interpretive line
drawing; the print comes from a right pes of a tyrannosaurid
theropod.
Cretaceous-Paleogene Boundary
at Amargos
Shallow-water Cretaceous-Paleogene bound-
ary sections in the Difunta Group, even though
potentially crucial as a link between regions of
extensive erosion on the coastal shelf and asso­
ciated deposition of reworked material in the
Gulf of Mexico, have not received much atten-
tion to date. Sedimentologically complex valley-­
like deposits in the La Popa area northwest of
Monterrey were interpreted by Lawton et al.
(2005) and Schulte et al. (2012) to be the result
of Chicxulub impact-induced tsunami backflow.
These channelized deposits were detected in the
Delgado Sandstone Member of the fluviatile to
shallow-marine Potrerillos Formation (Difunta
Group), which is considered to be coeval or
even a lithostratigraphic equivalent to the Las
Encinas Formation (e.g., Ifrim et al., 2010).
The main evidence for a near–Cretaceous-
Paleogene age of the deposits is reworked ejecta
of the Chicxulub impact (Lawton et al., 2005;
Schulte et al., 2012). This ejecta material is made
Geological Society of America Bulletin, v. 1XX, no. XX/XX 13
End-Maastrichtian tetrapod tracks from Mexico
IV
II metatarsal pad
up of vesicular spherules of altered glass, simi-
lar to spherules known from the deep-marine
Burgos Basin east and south of Monterrey and
there discussed to be of late Maastrichtian (e.g.,
Stinnesbeck et al., 2001) or Cretaceous-Paleo-
gene boundary age (e.g., Schulte et al., 2009).
However, these spherules are easily redepos-
ited into younger sediments and have variable
stratigraphic ages spanning from the late Maas-
trichtian to the early Danian. In Guatemala,
­Belize, and Haiti, they are generally reworked
and interbedded with early Danian sediments
of the Parvularugoglobigerina eugubina (Pla)
zone (Stinnesbeck et al., 1997, 2000; Keller
et al., 2001, 2003a, 2003b). Ocean Drilling Pro-
gram (ODP) Sites 1001 and 1049 are very con-
densed and incomplete. Unconformities span
from the early Danian to the late Maastrichtian
zone CF3 (~360–550 k.y.) in Blake Nose Sites
1049A, 1049C, and 1050C, and from P1a (1)
through zone CF4 (~2.9–3.43 m.y.) in Carib-
bean Sites 999B and 1001B. They therefore
cannot provide any evidence of deposition pre-
cisely at the Cretaceous-Paleogene boundary
(Keller et al., 1993, 2003a, 2013; Mateo et al.,
impact spherules is similar throughout the NW
Atlantic, Caribbean, and Central America up to
central Mexico and was therefore interpreted
to result from either current erosion correla-
tive with sea-level falls (Mateo et al., 2015), or
from large-scale platform failure caused by the
Chicxulub impact (Sanford et al., 2016).
The La Popa spherules are also reworked,
with the precise age of the spherule-bearing
sedi­ments unknown. Both Vega et al. (1999,
p. 108) and Lawton et al. (2001, p. 232) pointed
out that Cimomia haltomi (Aldrich, 1931), a
Paleo­cene nautiloid, is present in clasts (!) in
Mudstone Member, which was subsequently
interpreted as a Chicxulub impact-induced tsu-
nami deposit by Lawton et al. (2005). Clearly,
the presence of Paleocene nautiloids, which
evolved hundreds of thousands of years after the
Cretaceous-Paleogene boundary, would indicate
that the base of the Upper Mudstone Member
of the Potrerillos Formation is post–Cretaceous-
Paleogene in age, independent from the pres-
ence of Chicxulub spherules in the deposit.
A mixture of fossils of different shallow-
water facies in the deposit was interpreted by
­Lawton et al. (2005) and Schulte et al. (2009)
to be eroded from shoreward environments and
transported basinward by south- to southeast-­
directed turbulent, supercritical backflow of
runup surge(s) emplaced against the continent
by one or several tsunami(s). Lawton et al.
(2005) concluded that these violent offshore-
directed flows may have provided a mechanism
for transport of voluminous, ejecta-bearing
sediment and late Maastrichtian marine organ-
isms into deep-water Gulf of Mexico settings.
However, mixtures of shallow-water fossils are
also a characteristic feature of incised valley
fills that always mark a major sequence bound-
ary. Lawton et al. (2001) described a sequence 1
boundary for the contact between the Delgado
Sandstone and Upper Mudstone Member.
Stinnesbeck et al. (2005) therefore proposed that
the valley infill was a transgression conglomer-
ate at the base of the transgressive systems tract
(TST), as was also suggested by Lawton et al.
(2001) and Soegaard et al. (2003).
The identification of Chicxulub impact ejecta
(smectite spherules) at Amargos qualifies the
section as a Cretaceous-Paleogene boundary lo-
cality. It is the second site located in the ­Difunta
Group of northeastern Mexico and the first
site known from Coahuila. Amargos, although
genetically assigned to the Parras Basin (see
Vogt et al., 2016, their fig. 1), is located only
~27 km from the La Popa locality and is simi-
lar to this latter site with regard to the presence
 Stinnesbeck et al.

Sphenodiscus pleurisepta—
A The Last Ammonite in Mexico

The presence of well-preserved S. pleuri-

septa specimens in the spherule-rich siliciclastic
unit, even with spherules in the body chamber
(Fig. 7), provides unequivocal evidence that this
B ammonite persisted in the region to about the
Cretaceous-Paleogene boundary. However, the
last appearance of S. pleurisepta in the spherule-
bearing unit does not necessarily mean that this
ammonite was exterminated by the Chicxulub
impact, because the spherules in the unit must
have been reworked from older sediments and
therefore cannot represent the original air-fall
deposit. This is indicated by the abundance of
siliciclastic debris and oyster shells and rare-
ness of spherules, which can only be explained
by erosion, reworking, and transport from an
original nearshore environment. It is also sug-
gested by the fair preservation of spherules and
absence, in the deposit, of air-fall–indicative
characters such as angular to flaser-like shards,
welding, plastic deformation, or calcite cement,
and the abundance of reworked shallow-water
debris (e.g., Stinnesbeck et al., 2001; Keller
C et al., 2002, 2003a). Rather, the shallow-water
environment at Amargos was highly dynamic
and regularly reworked by storms and tides.
On the other hand, the excellent preservation of
S. pleurisepta specimens (with body chambers)
proves that these shells were not transported
over long distances. In that case, the taxon could
have survived the impact. Sphenodiscus pleuri-
septa was already known to be one of the last
Maastrichtian ammonites in the Americas and
the last taxon to exist in Mexico, reaching into
Figure 13. Footprints of azhdarchoid pterosaurs from the Rancho the uppermost Maastrichtian (Ifrim and Stinnes-
San Francisco site: (A) Manus print of a very large azhdarchoid beck, 2010; Stinnesbeck et al., 2012; Landman
pterosaur (wingspan ~8 m) and interpretive line drawing (right). et al., 2014, 2015). The presence of S. pleuri-
(B) Pes print of a medium-sized azhdarchoid pterosaur (wingspan septa in the spherule-rich siliciclastic deposit
~3–4 m). Below: Interpretive drawing of another pes print (same size at Amargos further extends the time range of
as B). (C) Anatomy of the pes of an azhdarchoid pterosaur from the this taxon to about the Cretaceous-Paleogene
early Late Cretaceous Crato Formation (NE Brazil; note the long boundary and, depending on the stratigraphic
metatarsus with respect to the length of the digits, the huge claws, position of the Amargos spherule unit, into the
and the webbing). earliest Paleogene. The survival of ammonites
across the Cretaceous-Paleogene-boundary has
also been discussed for other species (e.g., Jagt
of altered spherules in a channelized sandstone prior to the end of the Maastrichtian, but prior to et al., 2003; Machalski and Heinberg, 2005;
and microconglomeratic deposit. At Amargos, the Chicxulub impact. Landman et al., 2014, 2015).
the siliciclastic deposit occurs ~70 m above lay- The abundance of oysters forming both shell
ers containing well-preserved crustaceans (Vega hash of highly fragmented small individuals Amargos Spherule-Bearing Unit—
et al., 2013) dated to the latest Maastrichtian. and little-transported specimens larger than An Early Paleocene Deposit?
Its base is located only ~8.5 m above the unit 60 mm indicates that individual layers of the
containing trackways of birds, and pterosaurs silici­clastic Cretaceous-Paleogene boundary de- The spherule-bearing siliciclastic unit at
(and nonavian dinosaurs at San Francisco). This posit at Amargos represent distinct depositional Amar­gos may even be an early Paleocene de-
assemblage of tetrapod ichnotaxa represents an events, e.g., during a TST, and not a single tsu- posit (see above). In addition to the discussion
autochthonous fauna that must therefore have nami wash-up event. This is also suggested for presented already, this scenario is suggested by
existed during the latest Maastrichtian, only the siltstone beds devoid of pebbles intercalated the first appearance of small ostreoids assigned
a few thousands to tens of thousands of years between individual microconglomeratic layers. here to G. gorizdroae Vyalov, 1937, in sedi-

14 Geological Society of America Bulletin, v. 1XX, no. XX/XX

 End-Maastrichtian tetrapod tracks from Mexico

A
references), no report exists to present knowl-
edge about tetrapods from the Upper Maastrich-
tian (e.g., the Las Encinas Formation) in the
area. Only Schulte et al. (2012) reported on an
isolated dinosaur bone from a Chicxulub-spher-
ule–bearing unit of the Delgado Formation north
of Monterrey, but this specimen was evidently
reworked from older sediments and may even

B C
be a mosasaur. The fossil finds from the new
San Juan Amargos and Rancho San Francisco
localities now provide evidence for the presence
of avian and nonavian dinosaurs and pterosaurs
in the latest Maastrichtian, to only a few tens of
thousands of years prior to the Chicxulub im-
pact. This scenario agrees with data presented
D E by Lockley and Hunt (1995), who discovered
dinosaur tracks at only a few decimeters below
the Cretaceous-Paleogene boundary.
Avian theropods form the bulk of the ichno-
taxa and show a worldwide extremely rare Maas-
trichtian diversity, exceeding that of the Lance
Formation in Wyoming (Lockley et al., 2004).
Figure 14. Invertebrate trackway morphotypes from the Amar- This abundance and diversity of birds, combined
gos site near Paredón: (A) morphotype 1 (Cruziana type, prob- with the rarity of pterosaurs and dinosaurs, may
ably produced by a branchiopod or a cephalocarinid crustacean), indicate that these latter Mesozoic reptilians
(B) morphotype 2, (C) morphotype 3, (D) morphotype 4 (epibenthic were already in decline prior to the Cretaceous-
detritus feeder, e.g., Gastropoda and Annelida), and (E) morpho­ Paleogene boundary, as was previously sug-
type 5 (Ophiomorpha). gested by MacLeod et al. (1997) and Koeberl
and MacLeod (2002). However, the Amargos
and San Francisco beach deposits likely formed
ments directly underlying the channelized de- dominated by turkostreids (Vega et al., 2007). in a mud plain devoid of vegetation. Similar to
posit at Amargos, and within it. This taxon from We therefore hypothesize that this major change extant mud plains, this environment may have
the Paleogene of Central Asia (e.g., Afghani- in the oyster assemblages was caused by the been an unsuitable area for terrestrial carnivores
stan; Berizzi Quarto di Palo, 1970) was synony- Chicxulub impact. Cretaceous oysters from the but not for aerial ones, including pterosaurs.
mized with Ostrea (Turkostrea) turkestanensis American Gulf Coast area did not survive this First of all, mud plains are devoid of vegetation.
Romanovsky, 1878 by Stenzel (1971), but in event. Asian taxa, which were apparently less Any terrestrial predator can be easily spotted
the same volume (Treatise on Invertebrate Pa- affected by the Chicxulub impact, immigrated during early approach. Second, the mud-plain
leontology, Part N, Volume 3), three specimens and evolved in the coastal region of the west- sediments are rich in invertebrate life that is
were illustrated as Ostrea (Turkostrea?) goriz- ern Gulf of Mexico. The unit with Gorizdrella mostly accessible for pokers rather than inertial
droae Vyalov, 1937 (Stenzel, 1971, fig. J.115, gorizdroae is also identified at Rancho San feeders. Third, mud-plain sediments often have
p. 1143). In a later paper, Vyalov (1984) re- Francisco but not in association with spherules deep flow channels that may contain soupy or
peated his original specific assignation as Goriz- (Fig. 8). Similar turkostreid ostreids are found in quicksand-like sediments representing deadly
drella gorizdroae, to which we here refer. There the Paleogene of the Neuquén area of southern traps for large terrestrial vertebrates, while aerial
is thus some confusion regarding the specific as- Argentina (Griffin et al., 2005). The immigra- vertebrates can test the sediment before landing.
signation of this ostreid, but both Vyalov (1937, tion of Gorizdrella gorizdroae suggests that The organic-rich sediment of the mud plains was
1984) and Stenzel (1971) agreed that the taxon the Chicxulub impact did not have the global thus a favorable food source for birds and small
is best transferred to turkostreids, a group that extinction effect that has been assumed (e.g., edentulous pterosaurs that fed on small food
first appears in the Maastrichtian of Argentina Schulte et al., 2009). items (Langston, 1981; Lehman and Langston,
(Griffin et al., 2005) but thrived in the Paleogene 1996). However, for this kind of feeding, the
of the Northern Hemisphere. Evidence for Pre–Cretaceous-Paleogene substrate would have been very soft to allow
The presence of Central Asian ­turkostreids Boundary Changes in the Tetrapod the akinetic beak to open inside the substrate.
represents an extraordinary and dramatic change Communities Large azhdarchoids could also have foraged on
in the oyster assemblages with respect to earlier small birds and turtles, large epibenthic inverte-
Campanian and Maastrichtian oyster associa- Even though abundant and diverse dinosaur brates, and carcasses (Paul, 1987; Chatterjee and
tions of the Difunta Group. Those o­ yster popu- material has been discovered in the Campanian ­Templin, 2004; Witton, 2007; Witton and Naish,
lations were dominated by species of the Atlan- part of the Difunta Group by several paleontol- 2008), or caught fish in shallow waters (Padian,
tic and Gulf Coasts (e.g., Arctostrea, Exogyra, ogy work groups (e.g., Secretaria de Educación 1988), but they certainly were rare visitors. The
Flemingostrea, Pycnodonte), whereas Asian Pública de Coahuila, the Instituto de Geología contrasting abundance and diversity of avian and
ostreids were absent. In contrast, later Paleo- of the Universidad Nacional Autónoma de ptero­saurian tracks hint at a decline of Ptero­
gene associations from the Difunta Group are México, and by us, e.g., Vogt et al., 2016, for sauria before the end of the Maastrichtian.

Geological Society of America Bulletin, v. 1XX, no. XX/XX 15


Flora in Pre–Cretaceous-Paleogene
Boundary Sediment
Floral remains are concentrated in certain
a­ reas consisting of fine-grained sandstone of
buff to reddish color. The facts that the plant ma-
terial is unsorted, milled to pieces, and may com-
prise shattered trunks with a diameter of up to
0.6 m suggest a primary transport in high-­energy
flash floods. The accumulation of the plant de-
bris occurred in low-energy flow channels or
residual ponds. Some of the wood pieces show
different stages of decay, suggesting longer-
term floating or exposure to moist environment.
Rarely, remnants of very badly preserved leaves
and fragments of fructifications are found, which
either must have been part of the primary debris
flow, but were deposited toward the end of the
high-energy phase, or were blown in later. From
the state of preservation, it is difficult to assess
the taxa, but most of the wood may come from
coni­fers showing the typical fibrous texture. The
only halfway identifiable leaf we found resem-
bles that of Gingkoales with deeply lobed leaves
(e.g., Gingkoites australis from the Late Creta-
ceous of Australia).
CONCLUSIONS
Here, we presented two new sections of the
Las Encinas Formation (Difunta Group) near
Paredón in the Mexican state of Coahuila. Sedi-
ments at Amargos and San Francisco are latest
Maastrichtian to earliest Paleogene in age. We
discovered dinosaur, avian, and pterosaurian
tracks at the close distance of only 8.5 m be-
low a unit bearing altered spherules attributed to
the Chicxulub impact. The majority of tetrapod
trackways were produced by at least six different
types of birds. Much less abundant and uniform
trackways are from azhdarchoid pterosaurs.
One isolated manus print suggests a wingspan
of about 8 m. One single right footprint with a
length of ~200 mm was produced by a thero-
pod, probably a tyrannosaurid. This ichnofossil
assemblage was deposited during the last tens
to hundreds of thousands of years of the Maas-
trichtian. The unit containing altered smectite
spherules is interpreted as a channelized incised
valley or TST deposit; the unit may even have
been deposited during the early Paleocene, but
this needs to be substantiated by future studies.
The ammonite Sp. pleurisepta reaches into the
spherule-bearing unit and is thus the last ammo-
nite known from Mexico, reaching to the Creta-
ceous-Paleogene boundary, or possibly across.
The Amargos and San Francisco sediments
are rich in shallow-marine invertebrates (e.g.,
molluscs, arthropods), vertebrates (e.g., marine
turtles, sharks), and plant remains and thus pro-
16 Geological Society of America Bulletin, v. 1XX, no. XX/XX
Stinnesbeck et al.
vide a unique opportunity to evaluate a marginal
marine ecosystem turnover prior to the Creta-
ceous-Paleogene boundary, suggesting faunal
transitions in invertebrates and tetrapods as well
as the vegetation.
ACKNOWLEDGMENTS
We are grateful to Manuela Böhm, Fabio Hering,
Rafael Moreno, Maria Zimmermann, Javier González,
Alfredo di Stefano, Torrey Nyborg, and Hector ­Porras
for assistance during field work at Paredón, and to
Rodolfo Peña for providing access to his ranch at
San Francisco. Martin Lockley and one anonymous
reviewer, as well as GSA Bulletin Editors Brian Pratt
and Brad Singer are gratefully acknowledged for
their many helpful comments and corrections to this
manuscript. Financial support for this research was
provided by Deutsche Forschungsgemeinschaft (DFG
STI 128/30, to Stinnesbeck and Frey) and by Rolf
Gademann (Würzburg).
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Science Editor: Bradley S. Singer
Associate Editor: Brian R. Pratt
Manuscript Received 17 April 2016
Revised Manuscript Received 1 September 2016
Manuscript Accepted 1 October 2016
Printed in the USA